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Research Article
A new species of the Cyrtodactylus brevipalmatus group (Squamata, Gekkonidae) from the uplands of western Thailand
expand article infoL. Lee Grismer‡§, Attapol Rujirawan|, Siriwadee Chomdej, Chatmongkon Suwannapoom#, Siriporn Yodthong¤, Akrachai Aksornneam|, Anchalee Aowphol|
‡ La Sierra University, Riverside, United States of America
§ San Diego Natural History Museum, San Diego, United States of America
| Kasetsart University, Bangkok, Thailand
¶ Chiang Mai University, Chiang Mai, Thailand
# University of Phayao, Phayao, Thailand
¤ Thaksin University, Phattalung, Thailand

Corresponding author: Anchalee Aowphol ( fsciacl@ku.ac.th )

Academic editor: Thomas Ziegler
© 2023 L. Lee Grismer, Attapol Rujirawan, Siriwadee Chomdej, Chatmongkon Suwannapoom, Siriporn Yodthong, Akrachai Aksornneam, Anchalee Aowphol.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Grismer LL, Rujirawan A, Chomdej S, Suwannapoom C, Yodthong S, Aksornneam A, Aowphol A (2023) A new species of the Cyrtodactylus brevipalmatus group (Squamata, Gekkonidae) from the uplands of western Thailand. ZooKeys 1141: 93-118. https://doi.org/10.3897/zookeys.1141.97624
ZooBank: urn:lsid:zoobank.org:pub:34377D39-2C3B-4626-8C50-8335C128E462
Open Access

Abstract

An integrative systematic analysis recovered a new species of the Cyrtodactylus brevipalmatus group from the uplands of Thong Pha Phum National Park, Kanchanaburi Province in western Thailand. Cyrtodactylus thongphaphumensis sp. nov. is deeply embedded within the brevipalmatus group, bearing an uncorrected pairwise sequence divergence of 7.6–22.3% from all other species based on a 1,386 base pair segment of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2) and adjacent tRNAs. It is diagnosable from all other species in the brevipalmatus group by statistically significant mean differences in meristic and normalized morphometric characters as well as differences in categorical morphology. A multiple factor analysis recovered its unique and non-overlapping placement in morphospace as statistically significantly different from that of all other species in the brevipalmatus group. The description of this new species contributes to a growing body of literature underscoring the high degree of herpetological diversity and endemism across the sky-island archipelagos of upland montane tropical forest habitats in Thailand, which like all other upland tropical landscapes, are becoming some of the most imperiled ecosystems on the planet.

Keywords

Bent-toed gecko, genetics, Indochina, integrative taxonomy, montane forests, morphology

Introduction

The gekkonid genus Cyrtodactylus Gray, 1827 contains well over 350 named and unnamed species and constitutes the third largest vertebrate genus on the planet (Grismer et al. 2021a, b; Uetz et al. 2022). To date, its extensive distribution extends across at least eight biogeographic regions and crosses a number of well-established biogeographic barriers from South Asia to western Melanesia (Grismer et al. 2022a). The ecological plasticity, phylogenetic relationships, and geographic distribution among, and within its 32 geographically circumscribed monophyletic species groups, are indicative of its ability to disperse across ephemeral seaways, major river systems, basins, mountain ranges, and land bridges, followed by extensive in situ diversification within specific geographic areas (Grismer et al. 2020, 2021a, b, 2022a).

Within Indochina and northern Sundaland, the Cyrtodactylus brevipalmatus group is one of the most ecologically and morphologically specialized groups within Cyrtodactylus (sec. Grismer et al. 2020, 2021a, b). All members bear a similar morphology, behavior, and color pattern adapted to an arboreal life style (Grismer et al. 2022b). The latest phylogenetic taxonomic treatment of the group (Grismer et al. 2022c) described four new species from Thailand, resulting in ten described and potentially as many undescribed populations needing further study. One of these undescribed populations, C. sp. 9 from Thong Pha Phum National Park, Kanchanaburi Province in western Thailand (Fig. 1), was first recognized on the basis of molecular phylogenetic evidence from a single specimen (Chomdej et al. 2021). We collected and sequenced eight additional specimens which corroborate the results of Chomdej et al. (2021) in that all eight specimens plus the specimen of Chomdej et al. (2021) form a monophyletic lineage deeply nested within the brevipalmatus group (Grismer et al. 2022c). Univariate and multivariate analyses of the eight new specimens recovered statistically significant morphological and morphospatial differences from all other members of the group which unequivocally indicate that it requires species-level recognition (Grismer et al. 2022c). As such, it is described herein.

Figure 1. 

Distribution of nominal species and unnamed populations of the Cyrtodactylus brevipalmatus group. Stars denote type localities. White circles are literature localities from which specimens were not examined and remain unidentified. Locality data for all material examined is in Grismer et al. (2022c: table 1).

Materials and methods

Genetic data

Methods for DNA extraction, sequencing, and editing followed Grismer et al. (2021c) and resulted in a 1,386 base pair segment of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2) and adjacent tRNAs. All material examined is listed in Grismer et al. (2022c: table 1) along with GenBank accession numbers.

Morphological data

The morphological data taken included 17 meristic, 18 normalized morphometric, and eight categorical characters (Grismer et al. 2022c) (Table 1). Normalization of the morphometric characters followed the method of Chan and Grismer (2022).

Table 1.
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Descriptions of morphometric, meristic, and categorical characters.

Abbreviations Characters
Morphometric characters
SVL snout-vent length, taken from the tip of the snout to the vent
TL tail length, taken from the vent to the tip of the tail–original or partially regenerated
TW tail width, taken at the base of the tail immediately posterior to the postcloacal swelling
HumL humeral length, taken from the proximal end of the humerus at its insertion point in the glenoid fossa to the distal margin of the elbow while flexed 90°
ForL forearm length, taken on the ventral surface from the posterior margin of the elbow while flexed 90° to the inflection of the flexed wrist
FemL femur length, taken from the proximal end of the femur at its insertion point in the acetabulum to the distal margin of the knee while flexed 90°
TibL tibia length, taken on the ventral surface from the posterior margin of the knee while flexed 90° to the base of the heel
AG axilla to groin length, taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body
HL head length, the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout
HW head width, measured at the angle of the jaws
HD head depth, the maximum height of head measured from the occiput to base of the lower jaw posterior to the eyes
ED eye diameter, the greatest horizontal diameter of the eye-ball
EE eye to ear distance, measured from the anterior edge of the ear opening to the posterior edge of the bony orbit
ES eye to snout distance or snout length, measured from anteriormost margin of the bony orbit to the tip of snout
EN eye to nostril distance, measured from the anterior margin of the bony orbit to the posterior margin of the external nares
IO interorbital distance, measured between the dorsomedial-most edges of the bony orbits
IN internarial distance, measured between the external nares across the rostrum
EL ear length, greatest oblique length across the auditory meatus.
Meristic characters
SL supralabial scales, counted from the largest scale at the corner of the mouth or posterior to the eye, to the rostral scale
IL infralabial scales, counted from termination of enlarged scales at the corner of the mouth to the mental scale
PVT paravertebral tubercles between the limb insertions, counted in a straight line immediately left of the vertebral column
LRT longitudinal rows of body tubercles, counted transversely across the body midway between the limb insertions from one ventrolateral body fold to the other
VS longitudinal rows of ventral scales, counted transversely across the abdomen midway between limb insertions from one ventrolateral fold to the other
VSM transverse rows of ventral scales, counted along the midline of the body from the postmentals to just anterior to the cloacal opening, stopping where the scales become granular
TL4E expanded subdigital lamellae on the fourth toe proximal to the digital inflection, counted from the base of the first phalanx where it contacts the body of the foot to the largest scale on the digital inflection–the large contiguous scales on the palmar and plantar surfaces were not counted
TL4U small, generally unmodified subdigital lamellae distal to the digital inflection on the fourth toe, counted from the digital inflection to the claw including the claw sheath
TL4T total number of subdigital lamellae beneath the fourth toe, TL4E + TL4U = TL4T
FL4E number of expanded subdigital lamellae on the fourth finger proximal to the digital inflection, counted the same way as with TL4E
FL4U small generally unmodified subdigital lamellae distal to the digital inflection on the fourth finger, counted the same way as with TL4U
FL4T total number of subdigital lamellae beneath the fourth toe, FL4E + FL4U = FL4T
FS enlarged femoral scales, counted from each thigh and combined as a single metric
PCS enlarged precloacal scales, counted as a single metric
PP number of precloacal pores in males, counted as a single metric
FP femoral pores in males, counted from each thigh and combined as a single metric
BB number of dark body bands, counted from between the dark band on the nape and the hind limb insertions on the body
Categorical characters
FKT tubercles on the flanks (present or absent)
SC1 slightly enlarged medial subcaudals (present or absent)
SC2 single distinctly enlarged, unmodified, row of medial subcaudal scales (present or absent)
SC3 enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginated (yes or no)
DCT dorsolateral caudal tubercles (small or large)
VLF1 DCT forming a ventrolateral caudal fringe (narrow or wide)
VLF2 ventrolateral caudal fringe scales generally homogenous or not (yes or no)
TLcross cross-section of the tail (round or square)

Phylogenetic analyses

Following Grismer et al. (2022c), an input file implemented in BEAUti (Bayesian Evolutionary Analysis Utility) v. 2.4.6 was run in BEAST (Bayesian Evolutionary Analysis Sampling Trees) v. 2.4.6 (Drummond et al. 2012) on CIPRES (Cyberinfrastructure for Phylogenetic Research; Miller et al. 2010) in order to generate a BEAST phylogeny, employing a lognormal relaxed clock with unlinked site models and linked trees and clock models. bModelTest (Bouckaert and Drummond 2017), implemented in BEAST, was used to numerically integrate over the uncertainty of substitution models while simultaneously estimating phylogeny using Markov chain Monte Carlo (MCMC). MCMC chains were run using a Yule prior for 40,000,000 million generations and logged every 4,000 generations. The BEAST log file was visualized in Tracer v. 1.7.0 (Rambaut et al. 2018) to ensure effective sample sizes (ESS) were well-above 200 for all parameters. A maximum clade credibility tree using mean heights at the nodes was generated using TreeAnnotator v. 1.8.0 (Rambaut and Drummond 2013) with a burn-in of 1,000 trees (10%). Nodes with Bayesian posterior probabilities (BPP) of 0.95 and above were considered strongly supported (Huelsenbeck et al. 2001; Wilcox et al. 2002). Uncorrected pairwise sequence divergences were calculated in MEGA 11 (Tamura et al. 2021) using the complete deletion option to remove gaps and missing data from the alignment prior to analysis.

Statistical analyses

All statistical analyses were conducted using R Core Team (2018). A Levene’s test for the normalized morphometric and meristic characters was conducted to test for equal variances across all groups. Characters with equal variances (F ≥ 0.05) were analyzed by an analysis of variance (ANOVA) and TukeyHSD post hoc test. Those with unequal variances (F < 0.05) were subjected to Welch’s F-test and Games-Howell post hoc test.

Morphospatial clustering and positioning among the species was analyzed using multiple factor analysis (MFA) on a concatenated data set comprised of 38 characters including non-metric categorical characters which cannot be used in a principal component analysis (Suppl. material 1). The MFA was implemented using the mfa() command in the R package FactorMineR (Husson et al. 2017) and visualized using the Factoextra package (Kassambara and Mundt 2017). A non-parametric permutation multivariate analysis of variance (PERMANOVA) from the vegan package 2.5–3 in R (Oksanen et al. 2020) was used to determine the statistical significance of centroid locations and group clustering. The analysis used a Euclidean (dis)similarity matrix with 50,000 permutations based on the loadings of the first four dimensions recovered from the MFA. The highly morphologically derived Cytodactylus elok was not included so as to prevent biasing the morphospatial relationships among the other species (see Grismer et al. 2022b).

Results

Phylogenetic analysis

The BEAST analysis recovered the Thong Pha Phum population as being deeply embedded within the brevipalmatus group and the strongly supported (1.00) sister lineage to two sister groups composed of (1) C. interdigitalis, C. uthaiensis, and C. sp. 11 and (2) C. cf. ngati1, C. cf. ngati2, C. ngati3, C. ngati4, and C. ngati (Fig. 2). The uncorrected pairwise sequence divergence between the Thong Pha Phum population and all other species of the brevipalmatus group ranges from 7.6–22.3%. (Table 2).

Figure 2. 

Maximum clade credibility BEAST phylogeny of the Cyrtodactylus brevipalmatus group highlighting the new species described herein. Bayesian posterior probabilities (BPP) are listed at the nodes.

Table 2.
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Mean (minimum–maximum) percentages of uncorrected pairwise sequence divergence (p-distances) among the putative species of the Cyrtodactylus brevipalmatus group based on 1,386 base pairs of mitochondrial NADH dehydrogenase subunit 2 gene (ND2) and adjacent tRNAs. Intraspecific p-distance are in bold font. n/a = data not applicable.

Species 1. C. brevipalmatus 2. C. cf. ngati1 3. C. cf. ngati2 4. C. elok 5. C. fluvicavus 6. C. interdigitalis 7. C. kochangensis 8. C. ngati, C. ngati3 and C. ngati4 9. C. rivularis 10. C. rukhadeva 11. C. thongphaphumensis sp. nov 12. C. sp. 10 13. C. sp. 11 14. C. sp. 14 15. C. uthaiensis
N 1 1 1 1 7 1 1 7 2 2 9 1 1 1 1
1. n/a
2. 21.03 n/a
3. 21.68 4.39 n/a
4. 20.77 22.58 21.42 n/a
5. 18.86 (18.84–18.97) 10.64 (10.58–10.84) 11.02 (10.97–11.23) 20.15 (20.13–20.26) 0.10 (0.00–0.26)
6. 20.77 6.97 9.16 22.84 12.02 (12.00–12.13) n/a
7. 19.35 14.58 14.71 20.90 12.31 (12.26–12.31) 15.23 n/a
8. 20.70 (20.65–20.90) 3.30 (2.84–4.00) 3.71 (3.35–4.26) 21.11 (20.90–21.42) 11.34 (11.10–11.87) 8.13 (7.74–8.65) 14.58 (14.45–14.84) 0.84 (0.00–1.55)
9. 20.00 (19.74–20.26) 15.87 (15.61–16.13) 15.03 (14.84–15.23) 21.61 (21.42–21.81) 12.57 (12.26–13.03) 15.48 (15.23–15.74) 12.26 (12.00–12.52) 15.03 (14.71–15.48) 0.52
10. 20.65 (20.13–21.16) 15.42 14.84–16.00) 15.48 (14.84–16.13) 21.61 (21.16–22.06) 12.25 (11.61–13.03) 16.00 (15.35–16.65) 13.10 (12.52–13.68) 15.23 (14.19–16.23) 4.65 (3.61–5.68) 1.55
11. 20.34 (20.13–20.65) 7.93 (7.74–8.00) 9.51 (9.42–9.55) 22.02 (21.81–22.32) 9.75 (9.55–9.94) 8.96 (8.77–9.03) 13.22 (13.03–13.29) 8.81 (8.13–9.68) 13.12 (12.77–13.42) 13.25 (12.52–13.94) 0.22 (0.00–0.52)
12. 19.87 9.29 10.84 21.94 10.12 (10.06–10.32) 10.19 13.68 10.21 (10.06–10.45) 13.94 (13.68–14.19) 14.32 (13.68–14.97) 8.06 (7.87–8.13) n/a
13. 20.39 7.23 8.90 22.19 11.12 (11.10–11.23) 3.87 14.58 8.28 (8.00–8.65) 15.35 (15.10–15.61) 15.61 (14.97–16.26) 8.96 (8.77–9.03) 10.45 n/a
14. 6.45 20.90 20.65 20.00 18.34 (18.32–18.45) 20.13 19.10 20.52 (20.26–20.65) 19.74 (19.48–20.00) 20.00 (19.48–20.52) 19.60 (19.48–19.87) 18.84 19.61 n/a
15. 19.74 5.81 8.13 21.16 10.12 (10.06–10.32) 7.1 13.94 6.97 (6.58–7.61) 13.94 (13.68–14.19) 13.94 (13.29–14.58) 7.80 (7.61–7.87) 8.39 6.58 19.48 n/a

Statistical analyses

The ANOVA and TukeyHSD post hoc and Welch’s F-test and Games-Howell post hoc tests of the adjusted morphometric and meristic characters were consistent with the phylogenetic and pairwise distance data in recovering a number of statistically significant differences between the Thong Pha Phum population and all other species (Table 3). Thong Pha Phum population plotted separately in the MFA with meristic data contributing 16.5% of the inertia in dimension 1, categorical morphology contributing 15.3% of the inertia in dimension 2, and normalized morphometric data contributing 13.6% of the inertia in dimension 3 (Fig. 3). The PERMANOVA analysis recovered the morphospatial position of the Thong Pha Phum population as being statistically different from C. brevipalmatus, C. cf. ngati2, C. ngati3, C. ngati, C. fluvicavus, C. interdigitalis, C. rivularis, C. rukhadeva, and Cyrtodactylus sp. 13 (Table 4).

Figure 3. 

A MFA of the species-level lineages based on the BEAST phylogeny (Fig. 2) B Percent contributions of each data type to the inertia of dimensions 1–4 of the MFA. Percentage values on the bar graphs are the amounts of inertia for the respective dimensions.

Table 3.
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Significant p-values from the results of the ANOVA and Welch’s F (*) analyses comparing the normalized morphometric and meristic characters of Cyrtodactylus thongphaphumensis sp. nov. to other species of the Cyrtodactylus brevipalmatus group. Only species with and N > 2 are included. No significant differences were recovered for SVL. Abbreviations are in the Materials and methods.

Morphometric characters AG* HumL* ForL FemL TibL HL HW HD* ED* EE* ES EN* IO EL IN
C. brevipalmatus 0.01 < 0.001 < 0.001 0.03
C. fluvicavus 0.0 0.007 0.013 0.023 0.007
C. interdigitalis 0.00 0.007
C. ngati < 0.001 < 0.001 0.042 0.007 < 0.001 < 0.001 0.000
C. ngati3 0.001 0.01 0.03 0.003 0.043 0.001 0.019 0.019 0.003
C. rukhadeva 0.02 0.004 0.02 0.033
Meristic characters SL IL* PVT* LRT VS VSM TL4E TL4T FL4E FL4U* FL4T* FS PCS* BB*
C. brevipalmatus < 0.001 0.003 0.022 < 0.001 0.05
C. fluvicavus < 0.001 < 0.001 0.001 0.004 0.020
C. interdigitalis 0.003 < 0.001 0.005 0.043 0.01 < 0.001
C. ngati 0.003 0.016 0.011 0.000 < 0.001 < 0.001 < 0.001 < 0.001
C. ngati3 0.001 0.042 < 0.001 0.001
C. rukhadeva 0.029 < 0.001 0.002 0.001
Table 4.
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Summary statistics from the PERMANOVA analysis from the loadings of dimension 1–4 of the MFA comparing Cyrtodactylus thongphaphumensis sp. nov. to all other species the Cyrtodactylus brevipalmatus group with sample sizes > 1. Bold fonts denote significant differences.

OTU pairs F model R2 p-value p-adjusted
C. rukhadeva 88.504 0.847 0.000 0.001
C. cf. ngati2 56.471 0.876 0.020 1.000
C. ngati3 59.321 0.868 0.006 0.324
C. interdigitalis 85.773 0.896 0.002 0.112
C. ngati 134.367 0.937 0.006 0.332
C. brevipalmatus 80.229 0.879 0.001 0.025
C. fluvicavus 55.127 0.809 0.000 0.008
C. rivularis 9.485 0.542 0.022 1.000
C. sp. 13 30.716 0.793 0.022 1.000

Taxonomy

Given the phylogenetic delimitation of the Thong Pha Phum population (Fig. 2), its statistically significant diagnostic morphological differences (Table 3), its statistically significant diagnostic placement in morphospace (Fig. 3, Table 4), and its notable difference in pairwise sequence divergence from all other species (Table 2), we describe it below as new species.

Cyrtodactylus thongphaphumensis sp. nov.

https://zoobank.org/4BB0E9B3-1BFF-49BC-BF77-79BF8CC95D27
Suggested Common Name: Thong Pha Phum Bent-toed Gecko Figs 4, 5

Cyrtodactylus sp. 9 Chomdej et al. 2021: 2; Grismer et al. 2022b: 248; Grismer et al. 2022c: 115.

Type material

Holotype. Adult male ZMKU R 00953 from Thong Pha Phum National Park, Pilok Subdistrict, Thong Pha Phum District, Kanchanaburi Province, Thailand (14.69339°N, 98.40534°E, 914 m a.s.l.), collected by Korkhwan Termprayoon, Akrachai Aksornneam, Natee Ampai, and Siriporn Yodthong on 8 April 2019.

Paratypes. Adult males ZMKU R 00951, ZMKU R 00954 and ZMKU R 00956 and adult females ZMKU R 00950, ZMKU R 00952, ZMKU R 00955, and ZMKU R 00957 bear the same collection data as the holotype.

Diagnosis

Cyrtodactylus thongphaphumensis sp. nov. can be separated from all other species of the brevipalmatus group by the combination of having 12–14 supralabials, 8–10 infralabials, 30–36 paravertebral tubercles, 19–21 rows of longitudinally arranged tubercles, 30–34 longitudinal rows of ventrals, 150–173 transverse rows of ventrals, 8–10 expanded subdigital lamellae on the fourth toe, 11–14 unexpanded subdigital lamellae on the fourth toe, 20–24 total subdigital lamellae on the fourth toe; seven or eight expanded subdigital lamellae on the fourth finger, 10–12 unexpanded subdigital lamellae on the fourth finger, 18–20 total subdigital lamellae on the fourth finger; 12–16 total number of enlarged femoral scales, 12–16 total number of femoral pores in males; 15 precloacal pores in males; 15–17 enlarged precloacals; enlarged femorals and enlarged precloacals not continuous; proximal femorals smaller than distal femorals; small tubercles on forelimbs and flanks; large dorsolateral caudal tubercles and wide ventrolateral caudal fringe; ventrolateral caudal fringe composed scales of different size; tail square in cross-section; maximum SVL 76.6 mm; 3–5 dark transverse body bands (Table 5).

Table 5.
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Sex and raw meristic, categorical, and morphometric data used in the analyses from specimens in the Cyrtodactylus brevipalmatus group. Abbreviations: R/L = right/left; / = data unavailable.

Species Cyrtodactylus thongphaphumensis sp. nov. C. brevipalmatus C. cf. brevipalmatus (C. sp. 14) C. brevipalmatus
Institutional catalog number ZMKU R 00950 paratype ZMKU R 00951 paratype ZMKU R 00952 paratype ZMKU R 00953 holotype ZMKU R 00954 paratype ZMKU R 00955 pratype ZMKU R 00956 pratype ZMKU R 00957 pratype LSUHC 1899 LSUHC 15076 LSUHC 11788 THNHM 10670 THNHM 14112
Sex ♀ ♂ ♀ ♂ ♂ ♀ ♂ ♀ ♂ ♀ ♀ ♀ ♀
Meristic data
Supralabials (SL) 12 13 13 14 13 13 13 13 11 12 10 14 12
Infralabials (IL) 8 8 10 10 9 10 10 9 8 10 9 11 11
Paravertebral tubercles (PVT) 32 33 34 34 36 36 30 30 39 37 38 37 37
Longitudinal rows of tubercles (LRT) 21 19 20 20 21 21 19 19 15 16 17 16 14
Ventral scales (VS) 34 33 33 34 30 33 32 33 38 38 38 36 39
Ventral scales along middle of the body (VSM) 173 158 156 166 159 159 150 169 176 170 182 154 160
Expanded subdigital lamellae on 4th toe (TL4E) 9 10 9 8 10 8 9 9 7 8 9 8 8
Unmodified subdigital lamellae on 4th toe (TL4U) 12 14 13 12 13 12 11 13 13 11 11 11 12
Total subdigital lamellae 4th toe (TL4T) 21 24 22 20 23 20 20 22 20 19 20 19 20
Expanded subdigital lamellae on 4th finger (FL4E) 8 7 7 8 8 8 8 8 8 8 8 7 8
Unmodified subdigital lamellae on 4th finger (FL4U) 10 12 12 11 12 12 11 12 9 11 10 10 10
Total subdigital lamellae 4th finger (FL4T) 18 19 19 19 20 20 19 20 17 19 18 17 18
Enlarged femoral scales (R/L) 5R/7L 8R/8L 8R/8L 7R/8L 8R/8L 7R/8L 7R/6L 8R/8L 0 0 0 8R/8L 7R/7L
Total enlarged femoral scales (FS) 12 16 16 15 16 15 13 16 16 10 11 16 14
Total femoral pores in males (FP) / 16 / 14 15 / 12 / 7 / / / /
Enlarged precloacal scales (PCS) 17 15 15 15 15 15 15 15 7 7 7 8 7
Precloacal pores in males (PP) / 15 / 15 15 / 15 / 7 / / / /
Postcloacal tubercles (PCT) 2 2R/3L 3 3 2R/3L 2R/3L 3 2 3 3 2 3 3
Body bands (BB) 3 4 3 4 3 5 4 4 4 6 3 5 5
Species Cyrtodactylus thongphaphumensis sp. nov. C. brevipalmatus C. cf. brevipalmatus (C. sp. 14) C. brevipalmatus
Institutional catalog number ZMKU R 00950 paratype ZMKU R 00951 paratype ZMKU R 00952 paratype ZMKU R 00953 holotype ZMKU R 00954 paratype ZMKU R 00955 pratype ZMKU R 00956 pratype ZMKU R 00957 pratype LSUHC 1899 LSUHC 15076 LSUHC 11788 THNHM 10670 THNHM 14112
Sex ♀ ♂ ♀ ♂ ♂ ♀ ♂ ♀ ♂ ♀ ♀ ♀ ♀
Categorical data
Small tubercles on flank (FKT) present present present present present present present present present present present present present
Dorsolateral caudal tubercles (DCT) large large large large large large / large small small small / small
Ventrolateral caudal fringe narrow or wide (VLF1) wide wide wide wide wide wide / wide narrow narrow narrow / narrow
Ventrolateral caudal fringe scales generally homogenous (VLF2) no no no no no no / no no no no / no
Tail cross-section (TLcross) square square square square square square / square circular circular circular / circular
Slightly enlarged medial subcaudals (SC1) present present present present present present / present present present present / absent
Single enlarged medial subcaudal (SC2) absent absent absent absent absent absent / absent absent absent absent / absent
Enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) no no no no yes no / no no no no / no
Morphometric data
SVL 73.1 73.5 73.7 73.2 64.4 76.6 76.6 74.2 68.8 70.8 64.1 66.0 63.8
AG 34.8 33.9 35.4 33.6 28.5 37.1 33.2 35.1 35.7 33.4 30.1 30.0 26.5
HumL 8.4 7.2 9.0 9.0 7.2 8.0 8.1 8.6 9.7 9.3 8.0 9.6 9.5
ForL 9.5 9.1 9.2 9.8 9.2 10.0 8.6 9.8 9.9 9.8 8.9 8.2 8.7
FemL 12.8 11.6 12.3 12.5 10.9 13.7 10.8 12.5 12.0 12.6 11.5 11.7 9.8
TibL 10.5 10.1 10.6 10.6 9.9 11.1 10.0 11.4 11.6 12.2 10.5 9.7 8.2
HL 19.9 20.9 20.1 20.0 17.6 20.4 19.3 20.0 19.3 19.3 19.0 17.9 18.2
HW 14.5 14.3 15.7 13.9 12.8 14.7 14.4 14.1 13.2 13.8 12.3 12.3 12.0
HD 7.8 7.7 7.9 7.7 7.0 8.2 7.8 7.6 8.0 7.6 7.6 7.3 7.0
ED 5.0 5.1 5.0 5.0 4.8 5.6 5.3 4.9 5.2 4.5 4.3 5.3 4.4
EE 5.9 5.9 6.0 5.9 5.3 6.1 6.0 6.0 5.7 5.9 4.9 5.7 5.7
ES 7.9 8.5 7.9 7.9 7.3 8.2 7.9 7.9 7.4 7.6 7.0 7.0 7.2
EN 6.0 6.1 6.0 5.8 5.4 6.1 6.0 5.9 5.7 5.4 4.9 5.3 5.4
IO 5.4 5.5 5.8 5.5 4.9 5.7 5.6 5.3 5.4 4.7 4.7 4.2 5.2
EL 1.1 1.5 1.5 1.2 1.2 1.0 1.2 1.3 1.0 1.4 1.1 1.3 1.0
IN 2.3 2.4 2.2 2.0 2.0 2.3 2.2 2.2 1.7 2.1 2.3 2.1 2.2
Species C. elok C. fluvicavus C. interdigitalis
Institutional catalog number LSUHC 8238 LSUHC 12180 LSUHC 12181 ZMMU R-16144 ZMKU R 00959 ZMKU R 00958 ZMKU R 00960 ZMKU R 00961 ZMKU R 00962 ZMKU R 00963 ZMKU R 00964 THNHM 20226 paratype THNHM 20228 paratype
Sex ♀ ♂ ♂ ♀ ♂ ♂ ♂ ♀ ♀ ♀ ♀ ♀ ♀
Meristic data
Supralabials (SL) 11 8 13 9 12R/12L 13R/12L 13R/12L 11R/12L 12R/12L 13R/12L 12R/11L 14 12
Infralabials (IL) 11 8 11 9 10R/10L 10R/10L 9R/10L 10R/10L 10R/10L 10R/10L 10R/10L 9 8
Paravertebral tubercles (PVT) 0 0 0 0 30 28 27 27 28 26 28 32 33
Longitudinal rows of tubercles (LRT) 6 7 4 4 17 17 14 16 17 18 16 19 20
Ventral scales (VS) 45 45 47 36 34 37 33 30 36 37 39 42 40
Ventral scales along middle of the body (VSM) 190 225 234 192 155 154 155 172 164 175 170 187 170
Expanded subdigital lamellae on 4th toe (TL4E) 10 9 9 9 9R/9L 10R/10L 9R/9L 9R/9L 10R/11L 9R/10L 9R/9L 12 10
Unmodified subdigital lamellae on 4th toe (TL4U) 11 10 11 9 11R/11L 12R/11L 10R/10L 12R/12L 11R/11L 10R/10L 12R/13L 14 13
Total subdigital lamellae 4th toe (TL4T) 21 19 20 18 20R/20L 22R/21L 19R/19L 21R/21L 21R/22L 19R/20L 22R/22L 26 23
Expanded subdigital lamellae on 4th finger (FL4E) 8 9 9 9 8R/8L 8R/8L 8R/8L 8R/8L 7R/7L 8R/9L 7R/7L 9 8
Unmodified subdigital lamellae on 4th finger (FL4U) 12 13 9 8 10R/10L 10R/10L 10R/9L 11R/11L 10R/10L 9R/9L 10R/10L 12 11
Total subdigital lamellae 4th finger (FL4T) 20 22 18 17 18R/18L 18R/18L 18R/17L 19R/19L 17R/17L 17R/18L 17R/17L 21 21
Enlarged femoral scales (R/L) 0 0 0 0 5R/6L 4R/5L 5R/6L 6R/6L 5R/6L 5R/6L 6R/6L 11R/8L 10R/9L
Total enlarged femoral scales (FS) 0 0 0 0 11 9 11 12 11 11 12 14 19
Total femoral pores in males (FP) / 0 0 / 11 8 10 / / / / / /
Enlarged precloacal scales (PCS) 8 8 8 7 15 14 14 15 14 15 15 14 15
Precloacal pores in males (PP) / 8 8 / 15 14 14 / / / / / /
Postcloacal tubercles (PCT) 3 2 3 3 3R/2L 3R/2L 3R/3L 1R/1L 3R/2L 3R/3L 2R/2L 3 2
Body bands (BB) 5 5 3 3 3 3 3 3 3 3 3 5 5
Categorical data
Small tubercles on flank (FKT) absent absent absent absent present present present present present present present present present
Dorsolateral caudal tubercles (DCT) large large large large small small small small small small small small /
Ventrolateral caudal fringe narrow or wide (VLF1) wide wide wide wide narrow narrow narrow narrow narrow narrow narrow narrow /
Ventrolateral caudal fringe scales generally homogenous (VLF2) no no no no no no no no no no no yes yes
Tail cross-section (TLcross) square square square square circular circular circular circular circular circular circular circular /
Slightly enlarged medial subcaudals (SC1) absent absent absent absent present present present present present present present absent /
Species C. elok C. fluvicavus C. interdigitalis
Institutional catalog number LSUHC 8238 LSUHC 12180 LSUHC 12181 ZMMU R-16144 ZMKU R 00959 ZMKU R 00958 ZMKU R 00960 ZMKU R 00961 ZMKU R 00962 ZMKU R 00963 ZMKU R 00964 THNHM 20226 paratype THNHM 20228 paratype
Sex ♀ ♂ ♂ ♀ ♂ ♂ ♂ ♀ ♀ ♀ ♀ ♀ ♀
Single enlarged medial subcaudal (SC2) absent absent absent absent absent absent absent absent absent absent absent absent /
Enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) no no no no no no no no no no no yes /
Morphometric data
SVL 80.2 78.2 84.8 78.6 72.5 72.0 69.6 68.4 76.8 65.7 78.2 81.2 74.8
AG 39.7 37.8 41.5 36.2 33.4 33.6 32.0 30.4 35.6 30.6 38.1 34.5 33.7
HumL 10.2 9.1 10.1 1.7 9.1 8.8 9.0 8.0 10.0 7.5 10.1 9.8 10.2
ForL 11.5 11.7 11.8 10.2 10.5 10.3 10.5 10.1 11.1 8.8 10.8 10.6 10.5
FemL 12.9 14.2 14.6 13.1 13.1 12.5 12.5 13.5 14.1 11.5 13.9 14.7 13.2
TibL 13.5 14.0 13.8 12.3 11.3 10.6 10.2 9.9 11.2 9.4 12.3 13.1 11.9
HL 21.8 21.6 21.9 21.7 20.1 20.5 19.7 20.1 21.2 18.6 21.3 20.8 19.9
HW 15.6 16.1 15.9 15.1 14.0 13.4 12.9 13.0 14.9 13.0 15.4 14.0 13.4
HD 9.6 9.8 10.4 9.8 8.5 8.1 8.3 7.9 8.1 7.8 8.3 3.4 8.6
ED 4.8 5.0 5.7 5.0 5.0 5.0 4.9 4.7 5.1 4.5 5.3 5.3 5.5
EE 6.4 7.1 7.0 6.8 6.5 5.9 5.7 5.8 6.1 5.4 6.5 5.8 6.2
ES 8.6 8.7 9.5 8.6 8.5 8.3 8.2 8.1 9.2 7.3 9.3 8.3 7.8
EN 6.0 6.2 6.5 6.2 6.5 6.2 5.9 6.1 6.6 5.6 6.5 6.0 5.5
IO 5.7 5.4 5.4 3.9 5.5 5.4 5.3 5.1 5.6 5.0 5.6 4.8 4.7
EL 1.9 1.4 1.5 1.4 1.4 1.5 1.7 1.4 1.8 1.6 1.8 1.3 1.3
IN 2.7 2.6 2.5 3.1 2.3 2.4 2.5 2.3 2.3 2.3 2.6 2.1 2.2
Species C. kochangensis C. cf. kochangensis C. ngati C. ngati3 C. ngati4 C. cf. ngati1 C. cf. ngati2 C. rivularis
Institutional catalog number ZMKU R 00945 THNHM 01667 HNUE-R00111 IEBR 4829 VNUF R.2020.12 HNUE-R00112 FMNH 255454 FMNH 270493 FMNH 270492 FMNH 265806 NCSM 79472 ZMMU R-14917 NCSM 80100 ZMKU R 00947 ZMKU R 00946
Sex ♀ ♂ ♂ ♀ ♀ ♀ ♀ ♂ ♂ ♂ ♀ ♀ ♀ ♀ ♀
Meristic data
Supralabials (SL) 12R/13L 12 10 10 10 10 13 13 13 10 14 9 12 13R/12L 13R/12L
Infralabials (IL) 9R/9L 10 9 9 9 9 10 9 11 8 11 10 12 11R/10L 10R/9L
Paravertebral tubercles (PVT) 34 29 39 40 38 40 28 27 26 27 28 32 29 34 33
Longitudinal rows of tubercles (LRT) 14 19 18 18 17 22 19 18 17 19 18 24 19 20 18
Ventral scales (VS) 35 34 38 36 35 32 37 36 36 33 33 36 35 34 37
Ventral scales along middle of the body (VSM) 172 159 168 164 178 158 159 166 156 158 164 166 165 160 166
Species C. kochangensis C. cf. kochangensis C. ngati C. ngati3 C. ngati4 C. cf. ngati1 C. cf. ngati2 C. rivularis
Institutional catalog number ZMKU R 00945 THNHM 01667 HNUE-R00111 IEBR 4829 VNUF R.2020.12 HNUE-R00112 FMNH 255454 FMNH 270493 FMNH 270492 FMNH 265806 NCSM 79472 ZMMU R-14917 NCSM 80100 ZMKU R 00947 ZMKU R 00946
Sex ♀ ♂ ♂ ♀ ♀ ♀ ♀ ♂ ♂ ♂ ♀ ♀ ♀ ♀ ♀
Expanded subdigital lamellae on 4th toe (TL4E) 9R/8L 8 8 10 9 9 10 10 8 10 9 8 10 9R/9L 9R/9L
Unmodified subdigital lamellae on 4th toe (TL4U) 12R/11L 13 11 10 11 10 11 11 11 11 12 10 10 13R/13L 12R/13L
Total subdigital lamellae 4th toe (TL4T) 21R/19L 21 13 16 17 16 21 21 19 21 21 18 20 22R/22L 21R/22L
Expanded subdigital lamellae on 4th finger (FL4E) 8R/8L 8 6 6 7 6 8 8 8 8 9 7 9 8R/8L 8R/8L
Unmodified subdigital lamellae on 4th finger (FL4U) 10R/10L 12 9 9 9 9 10 10 10 10 8 9 10 11R/10L 12R/12L
Total subdigital lamellae 4th finger (FL4T) 18R/18L 20 15 15 18 15 18 18 18 18 17 16 19 19R/18L 20R/20L
Enlarged femoral scales (R/L) 6R/6L 7R/7L 10R/10L 9R/8L 10R/9L 8R/9L 9R/7L 8R/9L 9R/9L 8R/8L 9R/8L 7R/8L 7R/8L 8R/8L 6R/8L
Total enlarged femoral scales (FS) 12 14 20 17 19 17 16 17 18 16 17 15 15 16 14
Total femoral pores in males (FP) / 14 14 / / / / 14 15 13 / / / / /
Enlarged precloacal scales (PCS) 12 16 13 13 13 13 15 13 13 13 12 13 13 15 15
Precloacal pores in males (PP) / 16 / / / / 13 13 13 13 / / / / /
Postcloacal tubercles (PCT) 1R/1L 3 3 2 1 2 0 0 0 0 2 3 4 2R/2L 3R/3L
Body bands (BB) 5 5 6 6 6 6 3 4 3 3 3 3 3 3 4
Categorical data
Small tubercles on flank (FKT) present present present present present present present present present present present present present present present
Dorsolateral caudal tubercles (DCT) large large small small small small small small small small small small small large large
Ventrolateral caudal fringe narrow or wide (VLF1) wide wide narrow narrow narrow narrow narrow narrow narrow narrow narrow narrow narrow wide wide
Ventrolateral caudal fringe scales generally homogenous (VLF2) no no no no no no yes yes yes yes yes yes yes yes yes
Tail cross-section (TLcross) square / circular circular circular circular circular circular circular circular circular circular circular square square
Slightly enlarged medial subcaudals (SC1) present present present present present present / present present present present present present absent absent
Single enlarged medial subcaudal (SC2) absent absent absent absent absent absent / absent absent absent absent absent absent present present
Enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) no no no no no no / no no no no no no no no
Morphometric data
SVL 60.1 70.2 66.5 68.1 69.3 46.6 83.6 70.2 74.1 73.8 78.0 87.1 77.7 73.9 68.1
Species C. kochangensis C. cf. kochangensis C. ngati C. ngati3 C. ngati4 C. cf. ngati1 C. cf. ngati2 C. rivularis
Institutional catalog number ZMKU R 00945 THNHM 01667 HNUE-R00111 IEBR 4829 VNUF R.2020.12 HNUE-R00112 FMNH 255454 FMNH 270493 FMNH 270492 FMNH 265806 NCSM 79472 ZMMU R-14917 NCSM 80100 ZMKU R 00947 ZMKU R 00946
Sex ♀ ♂ ♂ ♀ ♀ ♀ ♀ ♂ ♂ ♂ ♀ ♀ ♀ ♀ ♀
AG 29.0 31.5 28.8 29.8 30.2 19.7 41.3 35.4 37.0 31.3 38.2 41.9 36.8 34.8 33.2
HumL 6.5 10.2 7.9 8.1 8.5 5.6 8.6 8.7 8.6 6.9 8.7 11.5 9.2 8.1 7.6
ForL 7.6 8.6 9.2 10.0 10.1 6.5 10.2 9.3 10.4 10.0 10.3 10.4 10.7 9.7 9.1
FemL 10.4 12.1 11.5 11.5 11.5 7.6 13.7 12.7 13.0 13.1 13.1 15.2 14.2 11.4 10.4
TibL 8.4 11.8 10.8 11.1 11.8 7.8 12.5 11.8 11.2 11.1 12.8 12.6 12.7 11.2 10.3
HL 17.3 18.3 20.1 20.4 20.7 16.1 21.7 20.6 20.3 20.7 21.2 22.1 21.4 20.3 19.3
HW 11.6 12.1 12.6 12.0 11.8 8.8 13.8 12.5 13.0 12.3 12.7 14.8 13.5 14.9 13.7
HD 6.5 7.8 7.4 7.2 6.6 5.1 9.2 8.4 9.1 7.6 8.3 8.7 9.2 8.2 8.2
ED 4.2 5.2 3.8 4.1 3.4 2.6 4.9 4.9 4.9 4.8 6.5 4.6 6.0 5.8 5.6
EE 5.0 4.9 5.8 5.5 5.9 4.4 6.9 6.1 6.2 5.7 5.3 6.5 6.2 6.5 6.2
ES 6.9 7.5 7.5 7.6 6.9 5.0 9.0 8.3 8.3 8.2 8.7 8.8 8.4 8.3 7.9
EN 5.2 5.5 6.7 6.3 6.2 4.5 6.5 6.2 6.1 6.2 6.2 6.6 6.0 6.1 5.8
IO 4.2 4.0 5.6 5.4 5.6 4.2 6.6 5.6 5.4 5.1 4.9 3.5 5.7 5.8 5.5
EL 1.0 1.3 0.8 0.8 0.7 0.3 1.3 1.1 1.2 1.0 1.5 1.2 0.9 1.1 1.1
IN 1.9 2.2 2.8 2.6 2.6 2.0 2.8 2.5 2.5 2.3 2.7 2.2 2.5 2.3 2.0
Species C. rukhadeva C. cf. rukhadeva C. sp. 11 C. sp. 13 C. sp. 13 C. uthaiensis
Institutional catalog number ZMMU R-16851 ZMMU R-16852 ZMKU R 00948 THNHM 24622 THNHM 24838 THNHM 03251 THNHM 03252 THNHM 03253 THNHM 03254 THNHM 01807 ZMMU R-16492 THNHM 00104 THNHM 27821 ZMKU R 00949
Sex ♂ ♀ ♀ ♂ ♀ ♂ ♂ ♀ ♂ ♂ ♂ ♀ ♀ ♂
Meristic data
Supralabials (SL) 11 9 14 11 13 13 11 12 13 12 11 12 15 13R/15L
Infralabials (IL) 10 11 9 10 10 10 10 10 11 10 9 10 11 10R/11L
Paravertebral tubercles (PVT) 27 30 30 26 28 27 27 30 30 26 30 33 29 33
Longitudinal rows of tubercles (LRT) 19 20 19 18 19 18 18 19 19 19 18 18 20 17
Ventral scales (VS) 34 43 38 38 36 37 37 39 34 35 34 37 36 36
Ventral scales along middle of the body (VSM) 154 152 165 162 158 157 159 168 160 161 160 159 165 159
Expanded subdigital lamellae on 4th toe (TL4E) 9 9 9 8 9 9 10 9 10 10 9 9 7 8R/(broken)L
Unmodified subdigital lamellae on 4th toe (TL4U) 11 11 12 11 13 12 12 15 13 13 10 12 12 12R/(broken)L
Total subdigital lamellae 4th toe (TL4T) 20 18 21 19 22 21 22 14 23 23 19 21 19 20
Expanded subdigital lamellae on 4th finger (FL4E) 9 8 8 7 8 8 8 8 8 8 10 8 8 7R/7L
Species C. rukhadeva C. cf. rukhadeva C. sp. 11 C. sp. 13 C. sp. 13 C. uthaiensis
Institutional catalog number ZMMU R-16851 ZMMU R-16852 ZMKU R 00948 THNHM 24622 THNHM 24838 THNHM 03251 THNHM 03252 THNHM 03253 THNHM 03254 THNHM 01807 ZMMU R-16492 THNHM 00104 THNHM 27821 ZMKU R 00949
Sex ♂ ♀ ♀ ♂ ♀ ♂ ♂ ♀ ♂ ♂ ♂ ♀ ♀ ♂
Unmodified subdigital lamellae on 4th finger (FL4U) 10 9 11 10 11 10 10 12 12 12 9 11 10 11R/11L
Total subdigital lamellae 4th finger (FL4T) 19 17 19 17 17 18 18 20 20 20 19 19 18 18R/18L
Enlarged femoral scales (R/L) 9R/8L 8R/8L 9R/8L 9R/L 9R/9L 9R/7L 7R/7L 6R/7L 5R/8L 7R/7L 9R/8L 9R/9L 7R/10L 8R/8L
Total enlarged femoral scales (FS) 17 16 17 18 18 16 14 13 13 14 17 18 17 16
Total femoral pores in males (FP) 17 / / 14 / 12 13 / 11 13 17 / / 12
Enlarged precloacal scales (PCS) 17 13 15 15 15 14 13 15 15 14 13 14 16 14
Precloacal pores in males (PP) 17 / / 15 / 14 13 / 15 14 13 / / 14
Postcloacal tubercles (PCT) 3 2 2R/3L 3 2 3 2 2 3 2 3 3 3 3R/3L
Body bands (BB) 3 3 3 3 3 4 4 / / 5 3 3 / 6
Categorical data
Small tubercles on flank (FKT) present present present present present present present present present present present present present present
Dorsolateral caudal tubercles (DCT) small small small small small small small small small / large small small large
Ventrolateral caudal fringe narrow or wide (VLF1) narrow narrow narrow narrow narrow narrow narrow narrow narrow / wide narrow narrow wide
Ventrolateral caudal fringe scales generally homogenous (VLF2) yes yes yes yes yes yes yes yes yes / yes yes yes no
Tail cross-section (TLcross) square square square square square square square square square / square circular circular circular
Slightly enlarged medial subcaudals (SC1) absent absent absent absent absent absent absent absent absent / present present present present
Single enlarged medial subcaudal (SC2) present present present present present present present present present / absent absent absent absent
Enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) no no no no no no no no no no no no no yes
Morphometric data
SVL 74.9 71.7 71.6 68.3 71.8 73.6 75.3 74.7 73.2 61.5 68.1 63.7 72.9 58.1
AG 34.6 32.6 33.9 27.3 29.9 30.9 31.3 32.2 30.3 26.2 34.6 25.8 30.6 26.6
HumL 10.7 10.4 7.9 9.8 8.3 12.2 11.3 11.8 11.0 10.1 10.3 7.6 10.1 7.0
ForL 8.6 7.9 9.6 8.7 8.5 9.0 10.6 9.6 9.2 7.9 8.5 8.1 9.6 8.3
FemL 12.6 11.8 10.5 10.8 10.9 11.5 10.2 11.9 12.1 9.5 12.6 10.7 12.8 10.0
TibL 10.1 9.3 11.2 9.7 10.7 10.9 11.7 11.3 11.1 9.1 11.4 10.1 10.2 8.4
HL 20.2 19.2 19.7 19.7 19.9 20.8 21.3 20.8 21.5 17.9 18.4 17.6 19.9 16.1
HW 14.6 13.4 14.0 13.1 13.9 14.9 15.0 13.1 14.1 11.8 13.1 11.9 13.8 10.9
Species C. rukhadeva C. cf. rukhadeva C. sp. 11 C. sp. 13 C. sp. 13 C. uthaiensis
Institutional catalog number ZMMU R-16851 ZMMU R-16852 ZMKU R 00948 THNHM 24622 THNHM 24838 THNHM 03251 THNHM 03252 THNHM 03253 THNHM 03254 THNHM 01807 ZMMU R-16492 THNHM 00104 THNHM 27821 ZMKU R 00949
Sex ♂ ♀ ♀ ♂ ♀ ♂ ♂ ♀ ♂ ♂ ♂ ♀ ♀ ♂
HD 9.2 8.5 8.3 7.3 8.9 8.2 8.2 8.1 8.9 7.5 8.3 7.7 8.4 6.3
ED 4.6 4.3 5.5 4.9 5.1 5.8 5.4 5.0 5.5 4.7 4.4 4.1 5.3 4.6
EE 6.2 6.2 5.8 5.1 6.2 5.6 5.7 5.4 6.2 4.3 6.2 4.9 6.3 4.7
ES 8.3 7.7 7.9 7.4 8.1 8.4 8.8 8.1 8.6 7.3 7.7 7.2 8.0 6.4
EN 6.3 5.7 5.8 5.4 6.0 6.2 6.4 5.8 6.2 5.3 5.5 5.6 5.9 4.9
IO 3.3 3.1 5.6 4.5 4.7 5.6 5.7 5.7 5.6 4.2 2.9 4.8 6.1 4.3
EL 1.2 1.0 1.4 1.6 1.5 1.2 1.3 1.2 1.2 0.9 0.9 1.4 1.4 1.5
IN 2.2 2.1 2.1 2.0 2.2 2.4 2.5 2.4 2.3 2.0 2.3 2.1 2.3 1.8

Description of holotype

(Fig. 4). Adult male SVL 73.2 mm; head moderate in length (HL/SVL 0.27), width (HW/HL 0.70), depth (HD/HL 0.39), distinct from neck, triangular in dorsal profile; lores concave slightly anteriorly, weakly inflated posteriorly; prefrontal region concave; canthus rostralis rounded; snout elongate (ES/HL 0.40), rounded in dorsal profile; eye large (ED/HL 0.25); ear opening horizontally elliptical, small; eye to ear distance greater than diameter of eye; rostral rectangular, divided by a dorsal furrow, bordered posteriorly by large left and right supranasals and one small azygous internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal, posteriorly by two unequally sized smaller postnasals, bordered ventrally by first supralabial; 14R/14L rectangular supralabials, second through eighth supralabials nearly same size as first, then tapering below eye; 10R/10L infralabials tapering smoothly to just below and slightly past posterior margin of eye; scales of rostrum and lores flat to slightly domed, larger than granular scales on top of head and occiput; scales of occiput intermixed with distinct, small tubercles; superciliaries subrectangular, largest anterodorsally; mental triangular, bordered laterally by first infralabials and posteriorly by large left and right trapezoidal postmentals contacting medially for 45% of their length posterior to mental; one row of enlarged, square to rectangular sublabials extending posteriorly to sixth(L) and fifth(R) infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth, imbricate, pectoral and ventral scales.

Figure 4. 

Adult male holotype of Cyrtodactylus thongphaphumensis sp. nov. ZMKU R 00953 (field no. AA 06933) from Thong Pha Phum National Park, Pilok Subdistrict, Thong Pha Phum District, Kanchanaburi Province, Thailand. A dorsal view B ventral view C dorsal view of head and ventral view of pelvic region D dorsal view of tail and E ventral view of tail in preservative F holotype in life.

Body relatively short (AG/SVL 0.46) with well-defined ventrolateral folds; dorsal scales small, granular interspersed with larger, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from occipital region onto base of tail and slightly beyond as paravertebral rows; smaller tubercles extend anteriorly onto nape and occiput, diminishing in size anteriorly; approximately 20 longitudinal rows of tubercles at midbody; approximately 34 paravertebral tubercles; tubercles on flanks; 34 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; 166 transverse rows of ventral scales; 15 large, pore-bearing, precloacal scales; no deep precloacal groove or depression; and two rows of enlarged post-precloacal scales on midline.

Forelimbs moderate in stature, relatively short (ForL/SVL 0.13); granular scales of forearm larger than those on body, interspersed with large flat tubercles; palmar scales rounded, slightly raised; digits well-developed, relatively short, inflected at basal interphalangeal joints; digits narrower distal to inflections; subdigital lamellae wide, transversely expanded proximal to joint inflections, narrower transverse lamellae distal to joint inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; 8R/8L expanded and 11R/11L unexpanded lamellae beneath the fourth finger; hind limbs larger and thicker than forelimbs, moderate in length (TibL/SVL 0.14), covered dorsally by granular scales interspersed with moderately sized, conical tubercles dorsally and posteriorly and anteriorly by flat, slightly larger, subimbricate scales; ventral scales of thigh flat, imbricate, larger than dorsals; subtibial scales flat, imbricate; one row of 6R/8L enlarged pore-bearing femoral scales not continuous with enlarged pore-bearing precloacal scales, terminating distally at knee; 7R/8L enlarged femoral scales; proximal femoral scales smaller than distal femorals, the former forming an abrupt union with much smaller, rounded, ventral scales of posteroventral margin of thigh; plantar scales flat, subimbricate; digits relatively long, well-developed, inflected at basal interphalangeal joints; 8R/8L wide, transversely expanded subdigital lamellae on fourth toe proximal to joint inflection extending onto sole, and 12R/12L unexpanded lamellae beneath the fourth toe distal to joint inflection; and claws well-developed, claw base sheathed by a dorsal and ventral scale.

Tail original, 94.6 mm long (TL/SVL 1.29), 5.0 mm in width at base, tapering to a point; nearly square in cross-section; dorsal scales flat, intermixed with tubercles forming paravertebral rows anteriorly and larger tubercles forming dorsolateral longitudinal rows; large, posteriorly directed, semi-spinose tubercles forming wide ventrolateral caudal fringe; larger scales of ventrolateral fringe occur at regular intervals; medial subcaudals enlarged but not paired, an enlarged single medial subcaudal longitudinal row absent; subcaudals, larger than dorsal caudals; base of tail bearing hemipenal swellings; 3R/3L conical postcloacal tubercles at base of hemipenal swellings; and postcloacal scales flat, imbricate.

Coloration in life

(Fig. 4). Ground color of the head body, limbs, and tail dull yellow; diffuse darker mottling on the top of the head; wider, pale-brown pre- and postorbital stripe extends from external nares to angle of jaw; whitish canthal and postorbital stripe dorsal to pale-brown pre- and postorbital stripe; faint, pale brown, nuchal band bearing two posteriorly directed projections; paired dark-brown paravertebral blotches on nape; four wide, irregularly shaped and broken transverse body bands edged in slightly pale brown between limb insertions; band interspaces bearing irregularly shaped scattered pale-brown markings; very faint pale-brown speckling on limbs and digits; seven wide pale-brown caudal bands separated by seven paler colored bands; posterior five pale-brown caudal bands encircle tail; ventral surfaces of body and limbs beige, generally immaculate, subcaudal region generally darker; iris orange-gold in color bearing black vermiculations.

Variation

(Fig. 5, Table 5). Individuals of the type series are very similar in overall coloration and pattern. TL and TW of complete original tails (ZMKU R 00951–00952, ZMKU R 00954, ZMKU R 00957) are 80.1–94.7 mm (mean 89.1 ± 6.5 mm; N = 4) and 4.2–4.9 mm (mean 4.7 ± 0.3; N = 4), respectively. ZMKU R 00956 has a short, partially regenerated tail which lacks banding (TL 27.7 mm, TW 5.1 mm). Similarly, the posterior sections of the tails in ZMKU R 00950 (TL 75.5 mm, TW 5.0 mm) and ZMKU R 00955 (TL 73.3 mm, TW 4.7 mm) are regenerated. Specimens ZMKU R 00950, ZMKU R 00952, and ZMKU R 00954 have three as opposed to four body bands in the holotype and ZMKU R 00955 has five body bands. Raw morphometric and meristic differences within and among all species of the brevipalmatus group are listed in Table 5.

Figure 5. 

Paratypes of Cyrtodactylus thongphaphumensis sp. nov. in preservative from Thong Pha Phum National Park, Pilok Subdistrict, Thong Pha Phum District, Kanchanaburi Province, Thailand.

Distribution

Cyrtodactylus thongphaphumensis sp. nov. is currently known only from the type locality at Thong Pha Phum National Park, Pilok Subdistrict, Thong Pha Phum District, Kanchanaburi Province, Thailand (Fig. 1).

Etymology

The specific epithet thongphaphumensis is in reference to the type locality of Thong Pha Phum National Park.

Comparisons

Cyrtodactylus thongphaphumensis sp. nov. is the sister species to a clade composed of eight lineages in the phylogenetic sequence of C. uthaiensis, sp. 11, C. interdigitalis, C. cf. ngati1, C. cf. ngati2, C. ngati3, and the sister lineages C. ngati4 and C. ngati (Fig. 2). Cyrtodactylus thongphaphumensis sp. nov. differs from those lineages by an uncorrected pairwise sequence divergence of 7.6–9.7% and from all members of the brevipalmatus group by 7.6–22.3% (Table 2). It differs discretely from C. elok by having as opposed to lacking paravertebral tubercles, femoral and precloacal pores, and by having 19–21 as opposed to 4–7 longitudinal rows of tubercles. It differs from C. brevipalmatus, C. fluvicavus, C. interdigitalis, C. ngati, C. ngati3, and C. rukhadeva in having statistically significant different mean values of combinations of the morphometric characters of AG, HumL, ForL, TibL, HL, HW, HD, EE, ES, EN, EL, and IN (Table 3). It differs further from those same species in having statistically significant different mean values of combinations of the meristic characters SL, PVT, LRT, VS, VSM, TL4T, FL4E, FL4U, FL4T, FS, PCS, and BB (Table 3). Discrete differences between Cyrtodactylus thongphaphumensis sp. nov. and other putative species and populations are presented in Table 5.

Natural history

All individuals were found in hill evergreen forest at 914 m elevation (Fig. 6). Specimens (N = 8) were collected at night (1900–2100 h) during the dry season (April) on tree trunks (62.5%; N = 5), on a building (12.5%; N = 1), and the ground (25.0%; N = 2) with a temperature of 27.0 °C and relative humidity of 71.1%. The holotype (ZMKU R 00953) and four paratypes (ZMKU R 00950, ZMKU R 00954, ZMKU R 00956–00957) were found on tree trunks ≤ 160 cm above ground level. One specimen (ZMKU R 00951) was found on a building. Two specimens (ZMKU R 00952, ZMKU R 00955) were found on ground. At night, the new species was found to co-occur with other gekkonid lizards, Cyrtodactylus oldhami (Theobald, 1876), Gekko kaengkrachanense (Sumontha, Pauwels, Kunya, Limlikhitaksorn, Ruksue, Taokratok, Ansermet & Chanhome, 2012), and Hemidactylus garnotii Duméril & Bibron, 1836.

Figure 6. 

Habitat of the type locality at Thong Pha Phum National Park, Pilok Subdistrict, Thong Pha Phum District, Kanchanaburi Province, Thailand.

Discussion

The discovery of new populations of the Cyrtodactylus brevipalmatus group across the archipelago of the upland sky-island habitats in Thailand will likely be commonplace with increased field work. Many such undescribed populations have already been reported and photographed on social networking platforms and these populations will be sampled and analyzed in order to ascertain their species status. Grismer et al. (2022c) pointed out that for several years many such populations went unanalyzed and were simply placed in the synonymy of either C. brevipalmatus or C. interdigitalis, only to be elevated later to species status following data-rich phylogenetic delimitation and morphological diagnostic analyses (Grismer et al. 2021c, 2022c). This current work not only contributes to an increased understanding of the unrealized diversity within the brevipalmatus group, but to a growing body of literature underscoring the high degree of herpetological diversity and endemism across a sky-island archipelago of upland montane tropical forests in Thailand (see Suwannapoom et al. 2022) which like many other upland tropical landscapes, are becoming some of the most imperiled ecosystems on the planet.

Acknowledgements

This work was financially supported by Office of the Permanent Secretary, Ministry of Higher Education, Science, Research and Innovation (Grant No. RGNS 64-038), Thailand Research Fund (DBG6080010) and Unit of Excellence 2023 on Biodiversity and Natural Resources Management, University of Phayao (FF66-UoE003). This research was reviewed and approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (ACKU61-SCI-008) and the Department of National Parks, Wildlife and Plant Conservation, Thailand provided the research permission. We would like to thank Charoen Jaichon (Thong Pha Phum National Park) for facilitating the fieldwork. Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Natee Ampai and Korkhwan Termprayoon assisted with fieldwork. Evan Quah, Vinh Luu, Olivier Pauwels, and an anonymous reviewer improved the manuscript.

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Supplementary material

Supplementary material 1 

Data frame for the multiple factor analysis of the putative species of the Cyrtodactylus brevipalmatus group

L. Lee Grismer, Attapol Rujirawan, Siriwadee Chomdej, Chatmongkon Suwannapoom, Siriporn Yodthong, Akrachai Aksornneam, Anchalee Aowphol

Data type: morphological data

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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